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Full Text in Pdf Format MARINE ECOLOGY PROGRESS SERIES Vol. 134: 119-130, 1996 Published April 25 Mar Ecol Prog Ser l Effects of macrobenthic burrows on infaunal communities in tropical tidal flats Sabine Dittmann* Zentrum fiir Marine Tropenokologie, Fahrenheitstr. 1, D-28359 Bremen, Germany ABSTRACT The effects of burrows on the infaunal community In a tropical t~dalflat were studied on the northeast coast of Australia. A comparative survey of infauna In burrows versus adjacent sediment was carr~edout for the burrows of 3 crustaceans (Scopimera inflata, Uca spp.. Callianassa austrahensis) and a brachiopod (Llngula anatjna). More (1.5 to 2.5~)infauna occurred within burrows than in adja- cent sediments Densities of meiobenthic nematodes, copepods and Platyhelminthes were significantly higher wlthin the burrows of Uca spp., C. australiensis and L. anatina. In the latter 2 cases multivariate analyses showed distinct communities in burrows and adjacent sediment, although this was not con- sistent over time for the brachiopod burrows. No differences in community composition were detected in the cases of Uca spp. and S. inflata. In a field experiment. C. austral~ensiswas excluded from 7 sites, and infaunal abundances and species compositions were followed for 1 yr and compared with control sites. From the very beginning, me~ofaunaldensities were significantly lower in the exclusion sites, mainly due to reduced densities of nematodes and copepods. The effect of the shrimp exclus~onon macrofauna was less pronounced, but after 1 yr total numbers of macrofauna were significantly lower in the exclusions, due to the distribution of amphipods. The results showed that promotive interactions play an important role in structuring tropical tldal flat communities. KEY WORDS: Benthos communities . Burrows . Callianassa . Field experiments Meiofauna . Promo- tlve interaction . Tropical tidal flat INTRODUCTION 1985, Branch & Pnngle 1987) The effects of biogenic structures have also been discussed in the light of Biogenic structures of benthlc organisms have been various functional-mode hypotheses (Rhoads & Young shown to have contrasting effects on infauna. On the 1970, Woodin 1976, Murphy 1985, Posey 1986, 1987, one hand, macrobenthic burrows may enhance the Dittmann 1990). presence of smaller infauna by providing suitable For tropical tidal flats, ambiguous accounts exist on microhabitats in sediment depth, where they would the relevance of biotic interactions for structuring otherwise not be able to live (Bell et al. 1978, Reise benthic communities. Sanders (1968) took tropical 1981, 1987, Meyers et al. 1987, Schaffner 1990). This shallow-water marine regions as an example of bio- process, termed accommodation, is one of the major logically accommodated communities, whereas Moore promotive interactions structuring benthic communi- (1972) and Alongi (1987, 1990) argued that the high ties (Reise 1985). Macrobenthos tubes can provide physical stress in tropical marine environments out- refuge from predation or facilitate larval settlement weighs biological processes in regulating intertidal (Woodin 1978, Bell & Woodin 1984, Bell 1985). On the populations. The existence of commensals associated other hand, infaunal abundances can be reduced in with intertidal macrobenthos has been reported from assemblages of burrowing organisms, an effect often the tropics (Kenway 1981, Morton & Morton 1983),yet attributed to bioturbation (Brenchley 1981, Murphy no quantitative study has been carried out to assess the - relevance of biogenic structures for the benthic com- 'Present E-mall [email protected]_wilhelmshaven.de munity in tropical tidal flats. 0 Inter-Research 1996 Resale of full art~clenot permitted Mar Ecol Prog Ser 134. 119-130, 1996 The aim of this investigation was to study whether MATERIALS AND METHODS burrows of macrobenthic organisms affect densities and/or species composition of smaller infauna in a The investigation was carried out between 1988 and tropical tidal flat. Decapod crustaceans are among the 1991 on the tropical northeast coast of Australia. Here, dominant macrobenthos dwellers in tropical tidal flats tidal flats extend along estuaries and bays and often (Rcise 1985, 1991, Alongi 1990, Dittmann 1995) and occur on the seaward side of mangrove forests. Tidal several burrowing crustaceans were chosen for this ranges are 2 to 2.5 m on average (Dittmann 1995). study. the sand bubbler crab Scopimera inflata Milne Assemblages of the brachiopod Lingula anatina and Edwards, 1873, fiddler crabs of the genus Uca, and the burrows of Scopimera inflata were studied in Hinchin- shrimp Callianassa australiensis (Dana). In addition, brook Channel; all other studies were carried out in dense patches of brachiopods (Lingula anatina La- the Haughton Estuary (Fig. 1). The sediment at the marck) were chosen as a further representative of studied sites consisted of fine sand (median grain size tropical macrobenthos. Densities of L. anatina on the 0.19 * 0.05 mm; sorting coefficient 1.59 + 0.34) with a Queensland coast, Australia, exceed 100 ind, m-' medium to low organic matter content [mud bank with (Kenchington & Hammond 1978). The hypothesis flddler crabs: 2.97% dry wt (DW);muddy sand flats: tested was that burrows increase infaunal abundances 1.38% DW]. and species numbers. To test this hypothesis, both a For a comparative survey, sediment samples were comparative study on infauna in burrows versus adja- taken from burrows of ocypodid crabs (Scopimera inflata cent sediment and an experiment in which C. aus- and Uca spp.), Lingula anatina and Callianassa aus- traliensis was excluded were carried out. traliensis, and compared with ambient sediment sam- The burrows of Callianassa australiensis extend over ples. These macrobenthic species occurred in different 1 m in depth and resin casts showed that the U-shaped communities (Dittrnann 1995, unpubl.). In each case, 5 or top part of the burrow merges at about 20 cm depth to 6 random samples were taken from burrows and the a single vertical tunnel that forms a branched network same number of replicates was taken from,the sediment of tunnels with horizontal chambers at over 50 cm surface adjacent (at least 1 cm distance) to burrows, all depth (Kenway 19811. On the surface, one of the open- within areas of 10 m2 The corers used for each burrow ings has a small cone-shaped mound c1 cm in eleva- type had a cross section just wide enough to include 2 to tion and is thus negligible in comparison to mounds of 5 mm of the sediment lining the respective burrow. Sam- other thalassinidean shrimps, which can reach eleva- ples were taken to a depth of 5 cm, and a lower horizon tions up to 1 m above the sediment surface (Griffis & (5 to 10 cm depth) was additionally sampled in burrows Suchanek 1991);the other opening has no mound. This of L. anatina to consider the effect of burrows on the burrow type represents a combination of burrow types vertical distribution of meiofauna in the sediment. 4 and 5 as classified by Griffis & Suchanek (1991). These authors specify shrimps of these types as filter/ suspension feeders, whereas Kenway (1981) describes the feeding process of C. australiensis as sand-sifting and showed that they feed while burrowing. This does not imply that material suspended while digging can- not be trapped on plumose setae on the antennae. inchinbrook In several temperate and subtropical locations, stud- ies have examined the effect of thalassinidean burrows on sediment reworking (Suchanek 1983, Suchan.ek & Colin 1986), microgeochemistry and microbial activi- ties (Aller et al. 1983, Dobbs & Guckert 1988) and infaunal abundances (Peterson 1977, Alongi 1986, Posey 1986, Branch & Pringle 1987, Posey et al. 1991). So far the studies have shown that the shrimp burrows provide a rich microbial and microalgal food source, but bioturbating activities exclude certain infauna spe- cies or functional modes. The results of the present investigation are discussed in relation to burrow type, effects of burrows reported from tidal flats elsewhere and previous accounts of communi.ty composition in beds of thalasslnidean Fig. 1. Northeast coast of Australia, show~ngthe location of shrimps. the study areas D~ttmannMacrobenthic burrows in trop~calt~dal flats Meiofauna was extracted by repeated shaking and layers of flyscreen (mesh size 1 5 mm) were implanted decantation through a set of sieves of 125, 80 and into the sediment at 7 of the sites before the sediment 40 pm mesh size. The samples were diluted with sea- was replaced in its oliginal position At the other 7 water, following narcotization with MgC1,. All sedi- sites, the sediment was ~eplacedwithout implanting ment samples were treated alive. screens and thus served as controls To aid in locating To study the effect of burrows of Callianassa aus- the sites, sticks were put into the sediment along the traliensis on community composition, an exclusion ex- site margins The arrangement of the sites was chosen periment was designed following the approach used haphazardly, making su~ethat the treatments were by Reise (1983) to exclude Arenlcola rnarlna from a inte~spersed Distances between sites were at least temperate tidal flat. C. australlensls is the structuring 2 m The flyscieen was effective in inhibiting the organism in the muddy sandflat of the studied inter- mobility of the shrlmps which could no longer reach tidal area (Dittmann 1995),and as ambient communi- the surface to irrigate their burrows and so moved ties differed in their sedimentological parameters, the away
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